Forensic Science International: Genetics 11 (2014) 111–116

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Forensic Science International: Genetics

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Forensic Population Genetics – Original Research

Analysis of 22 Y chromosomal STR haplotypes and Y haplogroup

distribution in Pathans of Pakistan
Eun Young Lee a,1, Kyoung-Jin Shin a,1, Allah Rakha b, Jeong Eun Sim a,
Myung Jin Park a, Na Young Kim a, Woo Ick Yang a, Hwan Young Lee a,*
a
Department of Forensic Medicine, Yonsei University College of Medicine, 50 Yonsei-ro, Seodaemun-gu, Seoul 120-752, Republic of Korea
b
Department of Forensic Sciences, University of Health Sciences, Lahore 54600, Pakistan

A R T I C L E I N F O A B S T R A C T

Article history: We analyzed haplotypes for 22 Y chromosomal STRs (Y-STRs), including 17 Yfiler loci (DYS19, DYS385a/
Received 6 August 2013 b, DYS389I/II, DYS390, DYS391, DYS392, DYS393, DYS437, DY438, DYS439, DYS448, DYS456, DYS458,
Received in revised form 3 March 2014 DYS635 and Y-GATA-H4) and five additional STRs (DYS388, DYS446, DYS447, DYS449 and DYS464), and
Accepted 4 March 2014
Y chromosomal haplogroup distribution in 270 unrelated individuals from the Pathans residing in the
Federally Administered Tribal Areas and the North-West Frontier Province of Pakistan using in-house
Keywords: multiplex PCR systems. Each Y-STR showed diversities ranging from 0.2506 to 0.8538, and the
Y chromosome
discriminatory capacity (DC) was 73.7% with 199 observed haplotypes using 17 Yfiler loci. By the
Y-STR
Haplotype
addition of 5 Y-STRs to the Yfiler system, the DC was increased to 85.2% while showing 230 observed
Haplogroup haplotypes. Among the additional 5 Y-STRs, DYS446, DYS447 and DYS449 were major contributors to
Pathans enhancing discrimination. In the analysis of molecular variance, the Pathans of this study showed
Pakistan significant differences from other Pathan populations as well as neighboring population sets. In Y-SNP
analysis, a total of 12 Y chromosomal haplogroups were observed and the most frequent haplogroup was
R1a1a with 49.3% frequency. To obtain insights on the origin of Pathans, the network analysis was
performed for the haplogroups G and Q observed from the Pathans and the Jewish population groups
including Ashkenazim and Sephardim, but little support for a Jewish origin could be found. In the present
study, we report Y-STR population data in Pathans of Pakistan, and we emphasize the need for adding
additional markers to the commonly used 17 Yfiler loci to achieve more improved discriminatory
capacity in a population with low genetic diversity.
ß 2014 Elsevier Ireland Ltd. All rights reserved.

1. Introduction obtained from many studies are accessible through an online

Y chromosomal short tandem repeats (Y-STRs) are informative
markers in investigations of sexual assault, paternity and genealog-
ical tests and evolutionary studies since they are inherited paternally
and their haplotype distribution may be distinct from other
populations of different geographic regions or with different ethnic
background [1–3]. In the field of forensics, a combination of 17 Y-STR
loci (DYS19, DYS385a/b, DYS389I/II, DYS390, DYS391, DYS392,
DYS393, DYS437, DYS438, DYS439, DYS448, DYS456, DYS458,
DYS635 and Y-GATA-H4), the so-called Yfiler haplotype, has been
commonly used to discriminate male individuals and to understand
the genetic structure of populations [4–6]. Y-STR haplotypes
* Corresponding author. Tel.: +82 2 2228 2482; fax: +82 2 362 0860.
E-mail addresses: hylee192@yuhs.ac, hylee192@gmail.com (H.Y. Lee).
1
These authors contributed equally to this work.
database, YHRD (Y chromosome haplotype reference database:
http://www.yhrd.org), which has reported 71,235 Yfiler haplotypes
as of December, 2013 (Release 46) [7]. The database shows that Yfiler
haplotype diversities may vary according to the genetic features of a
population; many populations have relatively high haplotype
diversity, but some populations that underwent a strong male
bottleneck in their history have low haplotype diversity [8–10].
Pathans are an Iranian-speaking Afghan ethnic group with a
widespread geographic distribution in southern and eastern parts
of Afghanistan and in north-west Khyber Pakhtunkhwa and
Baluchistan province of Pakistan [11,12]. Previous population
studies on this ethnic group have revealed relatively low Y
chromosomal haplotype diversity with 17 Yfiler loci [13,14]. Since
the lack of resolution may be a challenge in its forensic application
to male lineage discrimination as well as to individual identifica-
tion, additional Y-STRs should be added to increase discrimination
capacity in this population group.

http://dx.doi.org/10.1016/j.fsigen.2014.03.004
1872-4973/ß 2014 Elsevier Ireland Ltd. All rights reserved.
112 E.Y. Lee et al. / Forensic Science International: Genetics 11 (2014) 111–116

Here, we report Y chromosomal haplogroups and haplotypes for
22 Y-STRs including 17 Yfiler loci and 5 additional Y-STRs (DYS388,
DYS446, DYS447, DYS449 and DYS464) in 270 Pathan individuals
from Pakistan, and we investigate the relationship between the
Pathans and historically or geographically relevant populations.
2. Materials and methods
2.1. DNA samples
Blood samples were obtained from 270 unrelated male Pathans
residing in the North West Frontier Province (NWFP) and the
Federally Administered Tribal Areas (FATA) of Pakistan. All
participants gave their informed consent orally or in writing after
one of the authors explained the aims and procedures of the study,
which was approved by The Institutional Review Board of
Severance Hospital, Yonsei University in Seoul, Korea. DNA was
extracted using a QIAamp DNA mini kit (Qiagen, Hilden, Germany)
according to the manufacturer’s instructions.
2.2. Multiplex PCR and genotyping for 22 Y-STRs
To define genotypes of 22 Y-STRs, 17 Yfiler loci (DYS19,
DYS385a/b, DYS389I/II, DYS390, DYS391, DYS392, DYS393,
DYS437, DY438, DYS439, DYS448, DYS456, DYS458, DYS635 and
Y-GATA-H4) and 5 additional Y-STRs (DYS388, DYS446, DYS447,
DYS449 and DYS464) were amplified using three in-house
multiplex PCR systems as previously described [15]. The PCR
products were separated by capillary electrophoresis using an ABI
PRISM 310 genetic Analyzer (Applied Biosystems). A peak
detection threshold of 100 RFU was used for allele designation
using GeneMapper ID Software versions 3.2 (Applied Biosystems)
and each allele call at all STR loci was determined by comparing to
an allelic ladder. To ensure correct allele calls, Y-STR typing results
for more than 100 Pathan samples and the 9948 male control DNA
(Promega Corporations, Madison, MI, USA) were confirmed to be
comparable with the AmpFlSTR1 YfilerTM kit (Applied Biosystems,
Foster City, CA, USA).
2.3. Y chromosomal haplogroup determination
The 19 biallelic Y chromosomal markers (M40, RPS4Y711, M89,
M201, M69, M304, M172, M9, M20, M175, M122, M45, M242,
M207, P231, M17, M479, M124 and M184) were selected to
determine Y chromosomal haplogroups E, C, F, G, H, J, J2, K, L, O, O3,
P, Q, R, R1, R1a1a, R2, R2a and T, that are present in Eurasian
populations (Fig. 1). All but M172 marker that was analyzed by
direct sequencing were analyzed using in-house multiplex PCR and

Fig. 1. Phylogenetic tree of the 19 Y chromosomal binary polymorphisms analyzed in this study. The SNPs are indicated in each branch and the haplogroups are indicated at
the end of each branch according to Karafet et al. [38]. The number of haplotypes of each haplogroup was compared in two different combinations of Y-STRs.
 E.Y. Lee et al. / Forensic Science International: Genetics 11 (2014) 111–116 113

SBE reaction. The detailed methods for determination of Y
chromosomal haplogroup are described in the Supplementary
Material.
2.4. Statistical analysis
The following formulas were used: (1) discriminatory capaci-
ty = no. of different haplotypes/no. of individuals; (2)
P
locus or haplotype diversity ¼ ½n=ðn  1Þ  ð1  Pi2 , where Pi
is the frequency of the ith allele or haplotype and n indicates
the number of individuals [16]. To examine the relationship
between neighboring populations, pair-wise genetic distances
(Fst) and associated probability values (p-values, 10,000 permuta-
tions) were calculated using the analysis of molecular variance
(AMOVA), which is available on the YHRD website [7]. The Pathan
population of the present study was compared with the
Afghanistan Pathans (YHRD accession number YA003701), North
Afghanistan Pathans (YA003703), South Afghanistan Pathans
(YA003702), Afridi Pathans in India (YA003686), Yousafzai Pathans
in Pakistan (YA003748) and full population sets, including 483
haplotypes from Afghanistan [7,13,14], 2068 haplotypes from
India [7,10,17–19], 703 haplotypes from Iran [7,20] and 1089
haplotypes from Pakistan [7] based on the genotyping results for
15 Y-STR loci (DYS19, DYS389I/II, DYS390, DYS391, DYS392,
DYS393, DYS437, DY438, DYS439, DYS448, DYS456, DYS458,
DYS635 and Y-GATA-H4).
In addition, genealogical relationships among haplotypes of the
Pathan population and Jewish populations [21,22] within hap-
logroups G and Q were reconstructed using the program Network
4.6.1.1 [23]. Since all haplotypes overlap in only 8 Y-STR loci
(DYS19, DYS388, DYS389I/b, DYS390, DYS391, DYS392 and
DYS393), the 8 loci were used for the analysis, and the weighting
scheme took into account the Y-STR variation across the
haplogroup as described in a previous report [24]. AMOVA and
Fst analysis were also performed for haplotypes of Pathans and
Jewish populations using Arlequin software Version 3.5.1.2
[21,22,25].
3. Results
3.1. Diversity of 22 Y-STRs
We could successfully obtain genotypes of total 22 Y-STRs from
270 male individuals in a Pathan population except for one (PK-
164) which showed a null allele at two adjacent loci, DYS458 and
DYS449 (Supplementary Material Table S1), and the haplotype
data for 17 Yfiler loci has been deposited in the YHRD (Accession
number YA003846). The locus or haplotype diversity (h) and the
number of observed alleles or haplotypes for 19 single copy STR
loci and 2 multi-copy STRs are summarized in Supplementary
Material Tables S2 and S3. Among single copy loci, the DYS449
showed the highest diversity of 0.8339, and the DYS388 was the
least diverse locus (h = 0.2506) with seven different alleles. The 2
multi-copy Y-STRs, DYS385 and DYS464, were more diverse than
most of the single copy Y-STRs with diversity values of 0.8538 and
0.8258, respectively.
3.2. Haplotype resolution with additional Y-STRs
We evaluated the haplotype resolution of variously combined
Y-STRs (Table 1). The discriminatory capacity (DC) of the 17 Yfiler
loci was 73.7% with 199 different haplotypes. By the addition of 5
Y-STRs (DYS388, DYS446, DYS447, DYS449 and DYS464) to the 17
Yfiler loci, an improved discrimination capacity was obtained as a
DC of 85.2% from 230 observed haplotypes in 270 Pathan samples.
We investigated the change in haplotype diversity (h) and DC by
adding 5 Y-STRs to the Yfiler loci. In the case of adding one Y-STR to
the Yfiler loci, DYS449 showed the greatest increase in DC and h,
which were 78.5% and 0.9939, respectively. The improvement
achieved by DYS449 was followed by improvements with DYS446,
DYS464 and DYS447. Since DYS449 contributed the most to
increasing haplotype diversity, we then calculated the cumulative
DC and h after the addition of other Y-STR to 18 loci, including
Yfiler loci and DYS449. The DC value increased with the addition of
DYS446, DYS464 and DYS447, but did not change by the addition of
DYS388. When 3 loci were added to Yfiler loci, the combination of
DYS446, DYS447 and DYS449 was the most effective in increasing
the discrimination capacity. The combination of 5 additional Y-
STRs with the 17 Yfiler loci increased the h and DC to 0.9968 and
85.2%, respectively.
In addition, we examined the change in haplotype resolution
with two different combinations of Y-STR loci (Table 2). Using 17
Yfiler loci, only 169 haplotypes (62.6%) were observed once and the
most frequent haplotype was shared by 23 individuals. By adding 5
additional Y-STRs to the Yfiler loci, 211 haplotypes were observed
once, which accounted for 78.1% of the total observed haplotypes.
The most frequent haplotype was shared by 12 individuals, and
their unrelatedness was confirmed by the analysis of autosomal
STRs and mitochondrial DNA (mtDNA) control region sequence
(data not shown). The shared haplotypes by Yfiler loci were
separated to more distinct haplotypes by key markers, as indicated
in Supplementary Material Table S4. It was clear that the additional

Table 1
Number of haplotypes and haplotype diversities for each combined Y-STRs in 270 Pathan samples.

Haplotype Diversitya No. of haplotypes No. of unique haplotypesb Discriminatory capacity (%) Haplotype diversity

17 Yfiler loci 199 169 73.7 0.9903

Yfiler + DYS449 0.8339 212 188 78.5 0.9939
+DYS464 0.8258 210 185 77.8 0.9917
+DYS447 0.7714 203 175 75.2 0.9916
+DYS446 0.7672 211 185 78.1 0.9937
+DYS388 0.2506 200 171 74.1 0.9904
+DYS449, DYS446 222 200 82.2 0.9958
+DYS449, DYS464 219 198 81.1 0.9948
+DYS449, DYS447 217 196 80.4 0.9945
+DYS449, DYS388 212 188 78.5 0.9939
+DYS449, DYS446, DYS464 226 205 83.7 0.9964
+DYS449, DYS446, DYS447 227 208 84.1 0.9963
+DYS449, DYS447, DYS464 223 204 82.6 0.9954
+DYS449, DYS446, DYS464, DYS447 230 211 85.2 0.9968
22 Y-STRsc 230 211 85.2 0.9968
a
Locus or haplotype diversity of an added Y-STR.
b
A unique haplotype is defined as one that occurs only once in a given population according to [37] and the AmpFlSTR1 YfilerTM user’s manual.
c
22 Y-STRs: 17 Yfiler loci + DYS388, DYS446, DYS447, DYS449 and DYS464.
114 E.Y. Lee et al. / Forensic Science International: Genetics 11 (2014) 111–116
Table 2
Number of haplotypes observed in two different combinations of Y-STRs in 270
Pathan samples.
No. of haplotypes observed Combined Y-STRs
17 Yfiler loci 22 Y-STRs
n=1 169 (62.6%) 211 (78.1%)
n=2 22 13
n=3 3 2
n=5 1 3
n=6 1 –
n=7 2 –
n = 12 – 1
n = 23 1 –
Total 199 230
Y-STRs allowed for higher haplotype resolution. However, when
haplotypes were analyzed using all 22 Y-STRs together, DYS388 did
not affect the haplotype diversity because a haplotype separated
by DYS388 was also separated by DYS449.
3.3. Distribution of Y chromosomal haplogroups and haplotype
diversity within each haplogroup
In Y-SNP analysis, 12 different Y chromosomal haplogroups
were defined and the resultant haplogroup distribution was
mainly consistent with that of a previous study (Fig. 1 and
Supplementary Material Table S5) [26]. The R1a1a haplogroup
with South Asian origin was observed to be the most frequent
haplogroup, with 49.3% frequency. The G haplogroup was the next
most frequent with 14.1% frequency and was followed by the J2
(9.6%), R2a (7.0%) and L (5.9%) haplogroups. Every individual that
shared an identical haplotype belonged to the same haplogroup
when considering 17 Yfiler loci or 22 Y-STRs. Within the R1a1a
haplogroup, the DC was increased to 79.7% by using 22 Y-STRs in
comparison with 65.4% by Yfiler loci. DCs within 5 haplogroups, G,
L, Q, R2a and T were also enhanced by adding Y-STR markers to the
Yfiler loci. However, haplotype resolution of J2 haplogroup was not
changed by any additional Y-STR.
3.4. Population pair-wise genetic distances between neighboring
populations
In order to analyze the relationship between the Pathans of
Pakistan and the populations residing in a neighboring regions, our
population data was compared with those of neighboring Pathan
populations and several population sets available in the YHRD by
calculating pair-wise genetic distances. The analyses were
performed with 15 loci (Supplementary Material Table S6 and
Fig. S1) and there were significant differences between Pakistani
Pathans and other Pathan populations from neighboring regions
(p < 0.05). Moreover, Pakistani Pathans did not show similarity to
any other population sets from neighboring areas including
Afghanistan, India, Iran and Pakistan (p < 0.05).
3.5. Population relationship analysis to infer the origin of Pathans
In order to examine correlations of genetic structural back-
ground, genetic variation and distances were investigated between
the Pathan population of the present study and 2 Jewish
populations including Ashkenazim and Sephardim [21,22] based
on the haplotype data for 8 Y-STRs, DYS19, DYS388, DYS389I/b,
DYS390, DYS391, DYS392 and DYS393 (Supplementary Material
Tables S7 and S8). Of the total variance, 93.41% was resulted from
differences within groups, and 6.59% was attributable to differ-
ences among the 3 population groups. The Pathan population
showed significant differences from the Jewish population groups
and stayed away with similar distances.
In the median-joining network (MJ network) relating 49
haplotypes of the G haplogroup representing 106 Y chromosomes
(Supplementary Material Fig. S2), the Pathan and the Ashkenazi
Jewish populations had different dominant haplotypes from each
other (16–12–13–17–23–11–11–13 defined by alleles at loci in the
order DYS19, DYS388, DYS389I, DYS389b, DYS390, DYS391,
DYS392 and DYS393 for the Pathans and 15–12–14–18–23–10–
11–13 for the Ashkenazi Jews), and the Sephardic Jewish
haplotypes were dispersed sporadically without core haplotype.
A haplotype (15–12–13–17–23–10–11–13) was observed to be
shared by a Pathan and an Ashkenazi Jew, but this haplotype was
further differentiated by the addition of DYS439 (data not shown)
[21].
Within the MJ network of 16 haplotypes of the Q haplogroup
representing 41 Y chromosomes (Supplementary Material Fig. S3),
the haplotype which occupies the central position in the network
was 13–12–13–16–22–10–15–13, and it was shared by 20
chromosomes from 19 Ashkenazi Jews and a Sephardic Jew. A
haplotype (13–12–13–16–22–10–15–14) shared by a Pathan and 3
Ashkenazi Jews was also further divided into different haplotypes
by the addition of DYS439 (data not shown) [21].
4. Discussion
This study demonstrated the haplotype diversity of 22 Y-STRs
and its capacity to differentiate between individuals in the Pathan
population residing in Pakistan. Seventeen loci included in the
Yfiler system have been commonly used for haplotype analysis
because their discrimination capacity was sufficiently high in most
populations [4–6,8]. However, some population groups such as the
Pathans were reported to have relatively low haplotype diversity
with these loci, the reason for which was assumed to be the result
of cultural practices like patrilocal residence and polygyny [8–10].
With an increased number of analyzed Y-STRs, the higher
haplotype diversity can be obtained: therefore researchers have
examined various additional Y-STR loci to increase the haplotype
diversity and to improve the discriminatory capacity [8,27–29].
Therefore, in the present study, we evaluated the resolution
capacity of five Y-STRs (DYS388, DYS446, DYS447, DYS449 and
DYS464) in combination with 17 Yfiler loci.
DYS449 is considered a rapidly mutating (RM) Y-STR and has
excellent performance in male lineage discrimination in other
studies [29,30]. RM Y-STR loci are useful in forensic work due to
their high mutation rates, which are 6.5 fold higher than Yfiler STR
loci and allow differentiation between even close male relatives
[29]. This character of RM Y-STRs allows for more diverse
genotypes, particularly in the population groups with low
haplotype resolution such as in the Pathans. DYS446 with a
simple pentamer repeat motif did not stand out among various
markers in other populations [31]. However, DYS446 did contrib-
ute to the discrimination of more haplotypes in the Pathans despite
its relatively low gene diversity. This implies that haplotype
diversity is not only influenced by locus diversity of each combined
Y-STR. Moreover, the value of each Y-STR should be considered in
terms of its effect on haplotype discrimination when it is combined
with others because a marker can cause a different haplotype
distribution depending on the genetic structure of a given
population. The most polymorphic marker was DYS464 which
showed 46 different haplotypes in this study. Because DYS464 is a
multi-copy STR with four positions on the Y chromosome, it can
display various haplotypes and has high diversity. However, just
like at DYS385a/b loci, it is impossible to determine the order of
alleles without additional typing of each a, b, c and d loci in
DYS464, and thereby the diversity of DYS464 is usually
 E.Y. Lee et al. / Forensic Science International: Genetics 11 (2014) 111–116
underestimated. Nonetheless, DYS464 shows high diversity and
has been considered to be a useful discriminative marker, only that
the selection of DYS464 for forensic application must be made with
caution since the interpretation of profiles can be ambiguous when
it comes to male-male admixed specimens or degraded DNA
[32,33]. In the present study, because DYS464 showed a similar
pattern with DYS446 or DYS449 in differentiation of haplotypes,
the combination of DYS464 with DYS446 or DYS449 did not
increase haplotype diversity despite its high diversity and the
presence of various alleles. On the other hand, DYS447 showed a
different pattern from other loci and its addition to Yfiler loci
allowed for more diverse haplotypes in spite of its low gene
diversity.
After evaluation of the usefulness of additional markers, we
selected 3 Y-STR loci, DYS446, DYS447 and DYS449, that were
effective Y-STRs to increase discriminatory capacity in the Pathan
population, while DYS388 and DYS464 were excluded due to the
low discrimination capacity or possible difficulty in interpretation.
The low diversity of DYS388 in Pathans has also been reported in a
previous report [9]. The combination of these 3 Y-STRs with Yfiler
loci drew a DC of 84.1% which was greatly improved but still at a
lower level compared to other population data. Therefore, further
investigation into other markers is needed to achieve more
discrimination in the Pathans, and RM Y-STRs may be good
candidates since they present high differentiation capacity even in
the same male lineage [29,30]. When we compared population
differentiation, Pakistani Pathans showed a significant difference
from other Pathan populations as well as from neighboring
population sets, which suggests the need to construct their own Y-
STR database.
In Y-SNP analysis, the distribution of haplogroups showed a
pattern which was similar to those of previous studies in Pakistani
populations [26,34]. DYS388 alleles with 15 or more repeats were
restricted to the haplogroup J in the Pathans of the present study as
described in previous reports [24,35]. However, the haplogroup E
lineage, the presence of which can support the hypothesis on a
limited contribution of the Greeks to the Pathans [26], was not
found in this study. On the other hand, the Pathans also lay claim to
a possible Jewish origin [24]. Because haplogroups G-M201 and Q-
P36 have been suggested as minor Ashkenazi Jewish founding
lineages [21], the MJ network was constructed for the haplogroups
G and Q from the Pathans and the Ashkenazi and Sephardic Jewish
populations [21,22]. Parahaplogroup E-M35* and haplogroup J
(J2) are reportedly major Ashkenazi Jewish founding lineages
[21], but these lineages were not found in the Pathans of the
present study; the direct sequencing analysis of M172 marker
revealed that all of the Pathan samples belonging to the J
haplogroup are the J2 haplotypes. Because the R1a1a haplogroup
that is predominate in the Pathans as well as in European
populations was considered to have a complicated history like the
haplogroup J2 [21], these Y chromosomes were not used in the
comparison between Jewish and Pathan populations. In the MJ
network of the G haplogroup, only one haplotype was shared
between the Pathans and the Jewish populations, but this
haplotype was further divided into different haplotypes by the
addition of DYS439 [21]. In addition, the Pathan population formed
separate clades on the periphery of the network of the G
haplogroup with a different core haplotype from the Jewish
populations, which might suggest little contribution of the Jews to
the Pathans. In the MJ network of the Q haplogroup, a Pathan
sample shared a haplotype with 3 Ashkenazi Jews, and this
haplotype was also differentiated by the addition of DYS439 [21].
The absence of major Jewish founding lineages in the Pathans of
the present study and the rare haplotype sharing between the
Pathans and the Jewish populations suggest that there is little
support for a Jewish origin of the Pathans in the present study.
115
In addition, the high frequencies of the South Asian hap-
logroups, H, L, R1a1a and R2a (67.4%) and the low frequencies of
the Middle Eastern haplogroups G, J and T (25.2%) were not
consistent with the results of mtDNA study in the same population
group [36], where the high frequency of the West Eurasian lineages
(55.6%) and the relatively low frequency of the South Asian
lineages (39.1%) were observed. Therefore, further analyses of
other loci with more samples would be helpful to elucidate the
population history of the Pathans.
Conflict of interest
The authors declare that they have no conflict of interest.
Acknowledgements
This research was supported by the Future-based Technology
Development Program through the National Research Foundation
of Korea (NRF) funded by the Ministry of Education, Science and
Technology (nos. 2010-0020631 and 2011-0027729). The authors
would like to thank Mian Sahib Zar (Center of Excellence in
Molecular Biology, University of the Punjab, Pakistan) for his
valuable comments on the Pathans.
Appendix A. Supplementary data
Supplementary data associated with this article can be found, in
the online version, at doi:10.1016/j.fsigen.2014.03.004.
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